Programed cell death shapes the expression of horns within and between species of horned beetles.

Holometabolous insects provide an excellent opportunity to study both the properties of development as well as their evolution and diversification across taxa. Here we investigate the developmental basis and evolutionary diversification of secondary trait loss during development in the expression of beetle horns, a novel and highly diverse class of secondary sexual traits. In many species, horn growth during late larval development is followed by a period of dramatic remodeling during the pupal stage, including the complete resorption of horns in many cases. Here we show that programed cell death plays an important and dynamic role in the secondary resorption of pupal horn primordia during pupal development. Surprisingly, the degree of cell death mediated horn resorption depended on species, sex, and body region, suggesting the existence of regulatory mechanisms that can diversify quickly over short phylogenetic distances. More generally, our results illustrate that secondary, differential loss of structures during development can be a powerful mechanism for generating considerable morphological diversity both within and between species.